Interactive effects of temperature and predation on an estuarine zooplankton community

The winter planktonic copepod community of the estuaries near Beaufort N.C. underwent a consistent transition from dominance by Acartia tonsa Dana to dominance by Centropages spp., which was associated with unpredictable decreases in water temperature and increases in abundance of predatory mysids. Centropages subsequently remained dominant until spring. Experimental manipulations of water temperature and mysid abundance in enclosures showed that copepod species composition was determined by an interaction of direct temperature effects with predatory interactions among copepods and mysids. Low temperatures stimulated recruitment of Centropages spp. and inhibited recruitment of Acartia tonsa. Temperature decreases also apparently stimulated migration of mysids into the study area. While A. tonsa was dominant, selective predation by mysids reduced the relative abundance of A. tonsa, but after Centropages became dominant mysid predation had no effect on copepod species composition. Predation by Centropages on the nauplii of other species probably contributed to its persistent dominance in the estuary. Rising temperatures in the spring favored recruitment of Acartia tonsa and inhibited Centropages recruitment. Selective predation by fish entering the estuary in spring may have contributed to the decline in abundance of Centropages spp. and mysids.     

Predatory disturbance and prey species diversity: the case of gray whale (<Emphasis Type="Italic">Eschrichtius robustus</Emphasis>) foraging on a multi-species mysid (family <Emphasis Type="Italic">Mysidae</Emphasis>) community

Why competitive exclusion does not limit the number of coexisting plankton species is a persistent question for community ecology. One explanation, the intermediate disturbance hypothesis (IDH), proposes that elevated species diversity is a product of moderate levels of disturbance that allow the subsequent invasion of less competitive species. Here, we assess the shifts in species diversity in a mysid (Mysidae Dana, 1850) zooplankton community, where at least 10 species have, over the last 15 years, have come to comprise the primary prey base of summer resident gray whales (Eschrictius robustus Lilljeborg, 1861) in Clayoquot Sound, British Columbia. We evaluate trends in the community structure of mysids (species dominance, diversity, and richness) across mysid habitat in the study area during the gray whale foraging season (May–September) for the period 1996 and 2008. Mysid species composition varies among years and diversity has increased as whales shifted their predatory focus from benthic amphipods (Ampeliscidae Costa, 1857) to mysids, near our starting point in 1996. Holmesimysis sculpta Tattersall, 1933 was the dominant species in early years; however, in 2007, the dominance shifted to Neomysis rayi Murdoch, 1885. The habitat restrictions and life history attributes of local populations of coastal mysids leave them vulnerable to the cumulative impacts of increased predation pressure by gray whales. This case study presents a unique examination implicating predation as an agent of disturbance capable of altering the species structure of a local prey community.     

Experimental analysis of the predation impact of the larvae of pond smelt (<Emphasis Type="Italic">Hypomesus transpacificus nipponensis</Emphasis>) on zooplankton populations established in mesocosms

The predation impact of the larvae of pond smelt Hypomesus transpacificus nipponensis on a zooplankton community was studied using mesocosms. The fish significantly depressed the abundances of copepod nauplii and rotifers, especially Hexarthra mira. The vulnerabilities of these prey might be determined by their swimming behavior and population density, suggesting that larval fish selectively prey on zooplankton that have a high encounter rate with the predator. The larvae did not have a negative effect on the densities of cladocerans, but fish predation altered the cladoceran community structure from the dominance of B. longirostris to that of B. fatalis. This result suggests that larval fish predation is an important factor that shifts the species composition of Bosmina in some lakes, the shift occurring in the season when fish larvae are abundant. Our results have shown that predation by the larval fish would control not only the abundance, but also the community structure of the small-sized zooplankton prey.     

The possibility of predator avoidance by Lake Michigan zooplankton

Low densities of Diaptomus ashlandi, Diacyclops thomasi, and Daphnia galeata mendotae were measured at depths where Mysis relicta formed nighttime aggregations. Calculations suggest that mysid predation can not account for the rarity of prey animals at these depths, which further suggests that the prey avoided the mysids. Unlike D. galeata mendotae, Daphnia pulicaria was common in mysid aggregations. The somewhat larger size of D. pulicaria may reduce its vulnerability to mysid predation, and consequently may explain the vertical distribution differences between the two congeners. Vertical distributions of Limnocalanus macrurus and copepod nauplii showed no obvious relationships to the mysid distributions. These were the only two taxa with distributions that were correlated with chlorophyll a concentrations. All crustacean taxa were rare in the epilimnion at night when sonar recorded a dense fish school.     

Effects of chaetognath predation and nutrient enrichment on enclosed estuarine copepod communities

Summary The effects of predation by the chaetognath Sagitta hispida and nutrient enrichment on estuarine copepod community structure were studied by experimental manipulations in large enclosures. Chaetognath abundance and nutrient addition rates were manipulated in a factorial design. Predation by chaetognaths resulted in a significant decline in the relative abundance of Acartia tonsa and an increase in the relative abundance of Oithona colcarva. However, these effects were evident only at chaetognath densities far higher than observed in natural populations. Nutrient enrichment resulted in a decline in relative abundance of Paracalanus crassirostris, and, in the absence of chaetognath predation, an apparent increase in the relative abundance of A. tonsa. The effects of chaetognath predation were independent of enrichment level, apparently because of the absence of effects of enrichment on total copepod and chaetognath densities.     

The effect of prey composition and abundance on the predation rate and selectivity ofMysis relicta

The predation rate and selection of specific prey byMysis relicta in Lake Tahoe depends on both the total and the relative densities of prey classes. Functional responses of mysids to changes in the density of two species of calanoid copepod prey were curvilinear in laboratory experiments. In the field and in 2-prey treatments, preference was usually positive forEpischura and negative forDiaptomus. However, the preference forEpischura was greatest at the lowest combined densities of prey in the laboratory and decreased at densities >100Epischura / m3 in the field. Several hypotheses to explain the inverse relationship between selectivity and prey density are discussed. Field assemblages of prey vary vertically, horizontally and seasonally. Potential effects of this hetrogeneity on both predator and prey populations are examined.     

Factors affecting feeding selectivity of visual predators on the copepod Acartia tonsa: locomotion,visibility and escape responses

Visual predation by fish on copepods involves prey encounter, attack and capture; during any of these processes prey selection can occur. Developmental changes in copepods, including increases in swimming speed, size and image contrast increase the encounter rate and distance at which they can be detected by predators. Copepods compensate for this increase vulnerability with age through diel vertical migration and improved escape capabilities. This study quantifies the changes in swimming speed and movement pattern with developmental stage of the copepod Acartia tonsa, using a video-computer system for motion analysis. Changes in visible size and image contrast with developmental stage were quantified under simulated natural illumination conditions using a video based image analysis system. The escape responses of the naupliar stages of the copepod Acartia tonsa were quantified in response to a stationary pipette sucking in water at a constant speed. Accurate quantification of the parameters that affect feeding selectivity of planktivorous fish will provide the basis for evaluation of their relative importance in future studies.     

Impact of whitefish on an enclosure ecosystem in a shallow eutrophic lake: selective feeding of fish and predation effects on the zooplankton communities

Bag-type enclosures (75 m³) with bottom sheets and tube-type enclosures (105 m³) open to the bottom sediment were stocked with exotic whitefish (Coregonus lavaretus maraena) to study their predation effects on the plankton community. The fish fed mainly on adult chironomids during the period of their emergence (earlier part of the experimental period). Thereafter, the food preference was shifted to larvae of chironomids and crustacean zooplankters. The predation effects on the plankton community were not evident in the bag-type enclosures where zooplankton densities were consistently low. The fish reduced the crustacean populations composed ofBosmina fatalis, B. longirostris andCyclops vicinus in the tube-type enclosures where the prey density was high (above ca. 50 individuals 1⁻¹). The results suggested that the intensity of predation depended on the prey density. Rotifers increased in the fish enclosure, probably becauseCoregonus reduced the predation pressure byCyclops vicinus on rotifers and allowed the latter to increase. In the fish enclosures, no marked changes in species composition were observed. Zooplankton predated by the fish seemed to be distributed near the walls of the enclosures. Problems of enclosure experiments for examining the effects of fish predation on pelagic zooplankton communities are discussed.     

The changing ecosystem of a shallow, brackish lake, Hickling Broad, Norfolk, U.K. I. Trophic relationships with special reference to the role of Neomysis integer

• 1 Hickling Broad, a shallow, brackish lake in England, changed from dominance by submerged aquatic plants to dominance by phytoplankton in the 1970s. These changes were ascribed to the effects of guanotrophicatioh by black-headed gulls, and increased salinity which together resulted in fish kills caused by a toxic alga, Prymnesium parvum. A mysid, Neomysis integer, was believed to be important in switching the system to plankton dominance through its presumed selective feeding on Cladocera and increasing population size as fish predation decreased.
• 2 Studies on laboratory predation rates of the mysid on Cladocera and on the population dynamics, predation rates and diet of the mysid and the populations of the major zooplankter, Eurytemora affinis in the Broad in the 1980s, have shown that the former explanations were incomplete. The mysid could have markedly reduced the cladoceran community, for it has potentially high predation rates, but Cladocera were probably lost through salinization. Neomysis feeds efficiently on Eurytentora affinis, but the latter reproduces rapidly and its populations are unlikely to be controlled by mysid predation. Nauplii and small copepodites are selectively taken. Alternative and probably major food sources for the mysid are periphyton and benthic algae and detritus.
• 3 Roach and bream readily take the mysid but the present fish stock in the Broad is very low and can now exert minimal pressure on the mysid population. An invertebrate predator on the mysid, Palaemometes varians, is also unlikely to have major effects. The former model of the operation of the Broad's ecosystem needs reconsideration in view of the findings.

     

Different Predation Impacts of Two Cyclopoid Species on a Small-sized Zooplankton Community: An Experimental Analysis with Mesocosms

We used mesocosms to analyze predation impacts on the prey populations and prey community structures by two cyclopoid copepod species, the larger Mesocyclops pehpeiensis and the smaller Thermocyclops taihokuensis, who coexist with small-sized herbivorous zooplankton species in a fish-abundant lake. The overall predation impact on the prey populations was stronger for Mesocyclops than for Thermocyclops. Mesocyclops had a strong and less selective impact on the rotifer community but a selective impact on the crustaceans. In contrast, Thermocyclops had a selective predation impact on rotifers but a weak and less selective impact on the crustacean community. As a result, the former predator reduced the diversity of the crustacean community but not the rotifer community, while the latter had an opposite impact on the diversities of the two communities. It has been suggested that fish induce development of a zooplankton community dominated by the small-sized zooplankton species in fish-abundant lakes. Our results demonstrated that cyclopoid copepods altered species composition and diversity of the small-sized zooplankton community in such lakes. Thus, the results have given an important suggestion on the role of the invertebrate predator cyclopoid copepods, which often coexist with fish, that they determine population dynamics and community structures of small-sized zooplankton in fish-abundant lakes.     

The feeding rates ofPleurobrachia (ctenophora) andSagitta (chaetognatha), with notes on the potential seasonal role of planktonic predators in the dynamics of north sea zooplankton communities

The zooplankton off the north-east coast of England has been the subject of a number of studies focusing on its productivity. It has also been shown to be representative of the zooplankton of much of the western North Sea. The community contains a number of predatory species, three of which are widely described as ‘voracious’, the ctenophorePleurobrachia pileus, the chaetognathSagitta elegans and the hyperiid amphipodThemisto compressa (≡ Parathemisto gaudichaudi). This study investigates the role of these planktonic predators in this community, with special reference to the seasonal changes in predation pressure. The functional response ofPleurobrachia pileus feeding onAcartia was determined from laboratory experiments. It was found to be linear at prey densities typical of UK coastal waters, although the linear relationship appeared to break down at low and high prey densities. Feeding rate data forSagitta elegans were obtained from gut content analysis and published laboratory derived estimates of digestion time. Of the 1,789 individuals examined 198 (11.1%) had food remains in the gut. A linear relationship betweenSagitta length and prey size was established and the daily feeding rate ofSagitta elegans was estimated to be 0.4 prey items d⁻¹ ind⁻¹. For comparative purposes, the proportion of the copepod standing stock removed bySagitta elegans, Pleurobrachia pileus andThemisto gaudichaudi was estimated for each month of the year. From this model it was shown thatThemisto applied the most predation pressure, andSagitta elegans applied the least predation pressure of the three planktonic predators considered. The impact ofPleurobrachia will be to a large extent offset due to its peak of seasonal abundance coinciding with the zooplankton peak in the summer.     

Effect of prey size on vulnerability of copepods to predation by the scyphomedusae Aurelia aurita and Cyanea sp.

A growing number of studies correlate changes in zooplanktonpopulations with abundance of medusae, but we cannot yet explainor predict the specific factors driving these interactions.This study demonstrates that the size of copepods has a significantinfluence on their vulnerability to predation by scyphomedusae.This finding is important because prey size, independent ofbehavior, has been neglected in theoretical models of predationby medusae. In experiments in a planktonkreisel, we used liveand heat-killed prey (Acartia hudsonica adults and copepodites)to separate the effects of copepod size and behavior on feedingrates by two medusae (Aurelia aurita and Cyanea sp.). Resultsrevealed that: differences in copepod size had a significantimpact on feeding rates, and thus small size can provide a refugefrom predation; behavior of adults diminished the liabilityassociated with larger size; and medusae with different morphologiesingested A.hudsonica at similar rates. Other experiments demonstratedthat medusae digested copepods at different rates based on preysize and predator species, findings that have implications forall future laboratory and field studies that assess feedingby scyphomedusae. Finally, this study illustrates how laboratorystudies serve as critical supplements to field observations.The effect of prey size on feeding rates can be confounded bydifferences in prey behavior, yet explains why small copepodswere typically ingested at relatively low rates by medusae.Size was clearly a dominant factor influencing copepod vulnerabilityacross scyphomedusan species, even those with very differentmorphologies. Future work should focus on the mechanisms ofsize selection, or the factors influencing contact and retentionrates.     

Predator-prey interactions and community structure: chironomids,mosquitoes and copepods in Heliconia imbricata (Musaceae)

Summary Evidence from both field observations and experimental work indicates that predation by larvae of a midge, Pentaneura n. sp. (Chironomidae), causes the low densities of mosquito larvae (Culicidae) found in the water filled bracts of Heliconia imbricata (Musaceae), microhabitats typically colonized by mosquitoes. This predation affects 2 species of mosquitoes, Wyeomyia pseudopecten, a resident species, and Trichoprosopon digitatum, a non-resident species. Predation keeps resident mosquito densities low while completely excluding the nonresident mosquito from the habitat. Both these effects of predation depend on the presence of an abundant alternative prey, an undescribed species of harpacticoid copepod found in the bracts. These copepod prey sustain chironomids when resident mosquito densities are low, permiting predator densities to remain high enough to exclude the non-resident mosquito. I discuss the evolutionary and ecological implications of predation structuring communities.     

Trophic importance of the chaetognaths Eukrohnia hamata and Sagitta gazellae in the pelagic system of the Prince Edward Islands (Southern Ocean)

Trophodynamics and predation impact of the 2 dominant chaetognaths Eukrohnia hamata and Sagitta gazellae were investigated at 19 stations in the vicinity of the Prince Edward Islands and at a 24-h station occupied at the sub-Antarctic Front in late summer (April/May) 1996. During the entire investigation, the zooplankton assemblages were numerically dominated by copepods with densities ranging from 21 to 170 ind. m⁻³. Amongst the copepods, Clausocalanus brevipes, Metridia gerlachei and M. lucens dominated accounting for >90% of the total. Generally, chaetognaths were identified as the second most important group composing at times up to 30% (mean = 14.7%) of total zooplankton abundance. Of the two chaetognath species, E.␣hamata was generally numerically dominant. Gut content analysis showed that both chaetognath species are opportunistic predators generally feeding on the most abundant prey, copepods. No feeding patterns were evident during the 24-h station, suggesting that both species feed continuously. The feeding rates of E. hamata ranged from 0 to 0.50 prey ind. day⁻¹ and between 0 and 0.90 prey ind. day⁻¹ for S. gazellae. The maximum total predation impact of E. hamata was equivalent to 5.2% of the copepod standing stock or up to 103% of copepod production per day. For S. gazellae the predation impact was lower, reaching a level of 3.2% of the copepod standing stock or 63% of the daily copepod production. Chaetognaths can, therefore, be regarded as an important pelagic predator of the Prince Edward Islands subsystem. Received: 27 March 1997 / Accepted: 11 September 1997     

Induction of toxin production in dinoflagellates: the grazer makes a difference

The dinoflagellate Alexandrium minutum has previously been shown to produce paralytic shellfish toxins (PST) in response to waterborne cues from the copepod Acartia tonsa. In order to investigate if grazer-induced toxin production is a general or grazer-specific response of A. minutum to calanoid copepods, we exposed two strains of A. minutum to waterborne cues from three other species of calanoid copepods, Acartia clausi, Centropages typicus and Pseudocalanus sp. Both A. minutum strains responded to waterborne cues from Centropages and Acartia with significantly increased cell-specific toxicity. Waterborne cues from Centropages caused the strongest response in the A. minutum cells, with 5 to >20 times higher toxin concentrations compared to controls. In contrast, neither of the A. minutum strains responded with significantly increased toxicity to waterborne cues from Pseudocalanus. The absolute increase in PST content was proportional to the intrinsic toxicity of the different A. minutum strains that were used. The results show that grazer-induced PST production is a grazer-specific response in A. minutum, and its potential ecological importance will thus depend on the composition of the zooplankton community, as well as the intrinsic toxin-producing properties of the A. minutum population.     

Life histories ofNeomysis integer,and its copepod prey,Eurytemora affinis,in a eutrophic and brackish shallow lake

Data was collected on the population structure and fecundity of the mysidNeomysis integer and the calanoid copepodEurytemora affinis in Hickling Broad, a shallow and eutrophic brackish lake, over a two-year period 1988/89. Standing biomass and production rate estimates were made using estimates of size-specific dry weights and development times obtained from laboratory measurements, field observations and information found in the literature. Both mysid and copepod are capable of a rapid response to favourable conditions and have high rates of birth, growth and production.E. affinis reproduces throughout the year with an estimated annual (May 1988–May 1989) production of 20 g dry wt m⁻². Copepod standing biomass was less in 1989 compared with 1988 owing to an overall reduction in copepod body-size and a reduction of size at maturity. There was a suggestion from the data that this was caused by predation from a greater number of large (>9 mm body-length) mysids in 1989 compared with 1988.N. integer is highly seasonal in its growth with distinct peaks of recruitment in May and July. Annual (May 1988–May 1989) production of the mysid was estimated as 5.8 g dry wt m⁻². AlthoughE. affinis is the only available prey ofN. integer in the lake, the mysid population appears independent of changes in that of the copepod and probably avoids negative predator-prey feedback mechanisms owing to an ability to feed on epiphytic algae.     

Effect of light on the feeding rates of pelagic and littoral mysid shrimps: a trade-off between feeding success and predation avoidance

It has been shown that freshwater pelagic mysid shrimps capture zooplankton at a higher rate in light than in darkness. This has been suggested to be due to facilitation of visual predation on evasive zooplankton prey. To test this hypothesis with Baltic mysid shrimps, and to see whether pelagic (migrating) and littoral (non-migrating) mysids differ in this respect, we compared the feeding rates of Mysis mixta and Praunus flexuosus on the copepod Acartia spp. As light conditions change radically from the beginning of summer to late autumn at the Baltic latitudes, we conducted the experiments during three different times of the year to determine if there was a seasonal response to light in mysids. The feeding rates of pelagic mysids were significantly higher in total darkness than in light. In contrast, the feeding rates of littoral mysids did not differ in the dark and the light in the three different seasons. The decreased feeding of pelagic mysids under well-lit conditions may be an adaptation to avoid visual predation by pelagic fish. In contrast, littoral mysids, which live in the well-lit layer throughout the day, do not show a similar response. The fact that light did not increase feeding in either of the mysid species indicates that these mysid species do not use vision in capturing prey, but rely mainly on mechano-reception.     

Copepod growth and diatoms: insensitivity of <Emphasis Type="Italic">Acartia tonsa</Emphasis> to the composition of semi-natural plankton mixtures manipulated by silicon:nitrogen ratios in mesocosms

The feeding selectivity and the growth and reproductive success of the copepod Acartia tonsa have been studied in mesocosms fertilized at different Si:N ratios (0–1.75:1) and, therefore, at different compositions of the phytoplankton communities. Phytoplankton composition showed a strong response to nutrient ratios, with diatoms comprising >90% at Si:N ratios >1:1 of total biomass as opposed to <20% at the lowest ratio. A. tonsa strongly preferred feeding on motile prey (flagellates and ciliates) to feeding on diatoms. Nevertheless, diatoms comprised a substantial part of the diet at the highest Si:N ratios. A. tonsa egg production and the final (after 4 weeks) abundance of adults and copepodites showed no response to Si:N ratios while nauplii production slightly increased with Si:N ratios. It is concluded that the frequently reported deleterious effect of diatoms on copepod reproduction is rather unusual when copepods are confronted with a naturally diverse phytoplankton assemblage instead of clonal cultures in the laboratory.     

Food and growth relations of the marine microzooplankter,Synchaeta cecilia (Rotifera)

The trophic interactions of the marine rotifer Synchaeta cecilia were investigated by determining its feeding and growth rates on a wide variety of marine phytoplankton and by determining its susceptibility to predation by the calanoid copepod, Acartia tonsa. Reproduction of S. cecilia was sustained in four-day feeding trails by 13 of 37 algal species tested. Growth-supporting species included species of Cryptophyceae, Dinophyceae, Chlorophyceae and Haptophyceae in sizes from 4 to 47 μm. Within these taxa, other species in the acceptable size range failed to support growth. No species of Cyanophyceae, Bacillariophyceae, or Chrysophyceae supported growth of the rotifer. S. cecilia can be maintained on unialgal cultures of Cryptophyceae but growth is enhanced by a combination of two or three species; a mixture of Chroomonas salina (Cryptophyceae), Heterocapsa pygmaea (Dinophyceae), and Isochrysis galbana (Haptophyceae) has sustained laboratory stocks of S. cecilia for over four years. The expected response of S. cecilia to food quantity was observed: as food concentration was increased from 58 to 1154 μg C 1⁻¹, the population growth constant increased from 0.17 to 0.60 d⁻¹ at 20°C. This is equivalent to population doubling times of 4.0 and 1.1 days at H. pygmaea densities of 500 and 10⁴ cells ml⁻¹, respectively. The susceptibility of S. cecilia to predation was investigated by determining its rate of capture by the omnivorous marine copepod Acartia tonsa. At prey densities of 5 to 35 μg C 1⁻¹ (0.3 to 1.9 individuals 1⁻¹), A. tonsa readily ingested S. cecilia at rates up to 3 μg C copepod⁻¹ day⁻¹.     

The influence of prey behaviour on prey selection of the carnivorous plant Utricularia vulgaris

The mechanisms underlying differential prey selection of two microcrustaceans by the common bladderwort (Utricularia vulgaris) were studied in the laboratory. Functional response experiments with single prey showed that Utricularia had a higher attack rate coefficient and a longer handling time coefficient with the cladoceran Polyphemus pediculus than with the cyclopoid copepod Eucyclops serrulatus. Observation of predation rate, defined as number of prey eaten per unit time, from direct behavioural observation on single prey species, showed a higher predation rate with Polyphemus than on Eucyclops, at low prey densities. The opposite pattern was found at high prey density. When the two prey were presented simultaneously to the predator, Eucyclops was preferred over Polyphemus. Results from the situation with two prey and some of the results from the direct behavioural observations support field data on the diet of Utricularia, which shows that cyclopoid copepods are selected more frequently than Polyphemus.